Bucculatrix cristatella, copyright Ilia Ustyantsev.

Belongs within: Heteroneura.
Contains: Tineidae, Apoditrysia, Yponomeutoidea, Gracillariidae, Lyonetiidae, Psychinae.

The Ditrysia are a major clade of Lepidoptera accounting for some 98% of the species of the order. They are characterised by female genitalia with an internal connection between the copulatory opening and ovipore, as well as by large apodemes on the second abdominal sternite and short bands rather than long fibres in the proboscis muscles (Grimaldi & Engel 2005).

Basal members of the clade, historically gathered into the superfamily Tineoidea (now recognised as non-monophyletic) are small to medium-sized species with narrow, typically grey fore wings that often retain the M vein in the discal cell. Most lack ocelli though prominent ocelli are retained in the Douglasiidae (Nielsen & Common 1991). The Simaethistiidae are a small, poorly known group of basal ditrysians found in eastern Asia and Australia. The Psychidae are a diverse group with the proboscis vestigial or absent in adults (Nielsen & Common 1991). The Meessiinae are very small moths associated with fungi and lichens (Nielsen & Common 1991). The Roeslerstammiidae are characterised by the presence of two rows of scales on each antennal segment (Nielsen & Common 1991).

<==Ditrysia [Gracillarioidea, Papilionina, Tineina, Tineites, Tineoidea, Tortrices]
    |--Meessiinae [Meessiidae]KP19
    |    |--Meessia leopoldellaYS10
    |    |--DemobrotisNC91
    |    `--Eudarcia simulatricellaKP19
       |  |  `--Dryadaula [Dryadaulidae]KP19
       |  |       `--D. visaliellaKP19
       |  `--+--ApoditrysiaGE05
       |     `--+--+--YponomeutoideaKP19
       |        |  `--Roeslerstammiidae [Amphitheridae, Amphitherinae]NC91
       |        |       |--Amphithera heteroleucaKP19, NC91
       |        |       |--Enchoptila idiopisP27
       |        |       |--ThereutisNC91
       |        |       |--Macarangela leucochrysaNC91, C91
       |        |       |--NematobolaNC91
       |        |       |--Chalcoteuches phlogeraNC91
       |        |       `--RoeslerstammiaNC91
       |        `--+--GracillariidaeKP19
       |           `--+--BucculatricidaeNC91
       |              |    |--Ogmograptis scribulaNC91
       |              |    `--BucculatrixKP19
       |              |         |--B. alaternella Constant 1890C90
       |              |         |--B. cristatellaKP19
       |              |         |--B. gossypiiNC91
       |              |         |--B. helichrysella Constant 1890C90
       |              |         `--B. plataniRJ93
       |              `--+--LyonetiidaeKP19
       |                 |--BedelliaNC91 [BedelliidaeM03, Bedelliinae]
       |                 |    |--B. psamminellaL27
       |                 |    `--B. somnulentellaNC91
       |                 `--HeliodinidaeM03
       |                      |--Thylacosceles acridomimaP27
       |                      |--Heliodines princepsNC91
       |                      |--Epicroesa ambrosiaNC91
       |                      |--Baltonides roeselliformisRJ93
       |                      `--VanicelaP27
       |                           |--V. disjunctellaP27
       |                           `--V. xenadelphaP27
            |  i. s.: Liothula omnivoraWFS04
            |         Solenobia triquetrellaD51
            |         Iphierga macaristaNC91
            |         Epichnopteryx plumella (Denis & Schiffermüller 1775)JP05
            |         CanephoraJP05
            |           |--C. hirsuta Poda 1761JP05
            |           `--C. unicolorRD77
            |         Paramictoides febrettaYS10
            |         Pachythelia unicolorYS10
            |         ApteronaYS10
            |         PlutorectisG84
            |         CryptotheleaGE05
            |           |--C. cardiophagaIR05
            |           |--C. gloveriiVC07
            |           `--C. junodiF92
            |         LuffiaYS10
            |           |--L. ferchaultellaBC91
            |           `--L. lapidellaYS10
            |         Animula huebneriH01
            |         CochliothecaRD77
            |           |--C. crenulellaRD77
            |           `--C. helixRD77
            |         Acanthopsyche opacella [incl. A. atra]RD77
            |         AmictaRD77
            |         Sterrhopteryx pristinella Rebel 1934P92
            |    |--Thyridopteryx ephemeraeformisKP19
            |    `--OiketicusG84
            |         |--O. elongatusG84
            |         `--O. kirbyiL81
                 |--Narycia saxosaC70, P27
                 |--Trigonocyttara clandestinaC70
                 |--Lepidoscia arctiellaC70
                 |--Cebysa leucotelusC70, NC91
                      |--T. cawthronellaP27
                      `--T. tubulosa (Retzius 1783)JP05
Ditrysia incertae sedis:
  Azinis hilarellaWM66
  Cerostoma Latreille 1802L02
    |--*C. dorsatum [=Ypsolophus dorsatus]L02
    `--C. persicellaF92
  AcrolophusD01 [AcrolophidaeGE05]
    |--A. boucardi Druce 1901D01
    |--A. linus Druce 1901D01
    `--A. underwoodi Druce 1901D01
  Eucryptogona [Eriocottidae]NC91
    `--E. trichobathraNC91
    |--Douglasia Stainton 1854 [=Douglassia (l. c.) non Bartsch 1934]P66
    |--Tinagma leucanthesNC91
  Melusina [Compsoctenidae]RD77
    `--M. energaRD77
  Euplocamus guttellaG20

*Type species of generic name indicated


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[B37] Balduf, W. V. 1937. Bionomic notes on the common bagworm, Thyridopteryx ephemeraeformis Haw., (Lepid., Psychidae) and its insect enemies (Hym., Lepid.). Proceedings of the Entomological Society of Washington 39 (7): 169–184.

[C70] Common, I. F. B. 1970. Lepidoptera (moths and butterflies). In: CSIRO. The Insects of Australia: A textbook for students and research workers pp. 765–866. Melbourne University Press.

[C90] Constant, A. 1890. Description de dix espèces nouvelles de microlépidoptères de la faune française. Annales de la Société Entomologique de France, 6e série 9: cxxiv–cxxvi.

[C91] CSIRO. 1991. The Insects of Australia: A textbook for students and research workers 2nd ed. vol. 1. Melbourne University Press: Carlton (Victoria).

[D51] Dobzhansky, T. 1951. Genetics and the Origin of Species 3rd ed. Columbia University Press: New York.

[D01] Druce, H. 1901. Descriptions of some new species of Lepidoptera from east Africa and tropical America. Annals and Magazine of Natural History, series 7, 7: 432–444.

[F92] Fan Z. 1992. Key to the Common Flies of China 2nd ed. Science Press: Beijing.

[G84] Gauld, I. D. 1984. An Introduction to the Ichneumonidae of Australia. British Museum (Natural History).

[G20] Goldfuss, G. A. 1820. Handbuch der Naturgeschichte vol. 3. Handbuch der Zoologie pt 1. Johann Leonhard Schrag: Nürnberg.

[GE05] Grimaldi, D., & M. S. Engel. 2005. Evolution of the Insects. Cambridge University Press: New York.

[H01] Hampson, G. F. 1901. The Lepidoptera-Phalaenae of the Bahamas. Annals and Magazine of Natural History, series 7, 7: 246–261.

[IR05] Idris, A. B., & A. Rizki. 2005. Notes on the tribe Ephialtini (Hymenoptera: Ichneumonidae: Pimplinae) of Malaysia. Serangga 10 (1–2): 111–126.

[JP05] Jungmann, E., & U. Poller. 2005. Neunachweise und Wiederfunde für die Schmetterlingsfauna (Insecta; Lepidoptera) des Altenburger Landes. Mauritiana 19 (2): 317–326.

[KP19] Kawahara, A. Y., D. Plotkin, M. Espeland, K. Meusemann, E. F. A. Toussaint, A. Donath, F. Gimnich, P. B. Frandsen, A. Zwick, M. dos Reis, J. R. Barber, R. S. Peters, S. Liu, X. Zhou, C. Mayer, L. Podsiadlowski, C. Storer, J. E. Yack, B. Misof & J. W. Breinholt. 2019. Phylogenomics reveals the evolutionary timing and pattern of butterflies and moths. Proceedings of the National Academy of Sciences of the USA 116 (45): 22657–22663.

[L02] Latreille, P. A. 1802. Histoire Naturelle, générale et particulière des crustacés et des insectes vol. 3. Familles naturelles des genres. F. Dufart: Paris.

[L27] Lindsay, S. 1927. A list of the Lepidoptera of Deans Bush, Riccarton, Canterbury. Transactions and Proceedings of the New Zealand Institute 57: 693–696.

[L81] Long, J. L. 1981. Introduced Birds of the World: The worldwide history, distribution and influence of birds introduced to new environments. Reed: Sydney.

[M03] McQuillan, P. B. 2003. The giant Tasmanian ‘pandani’ moth Proditrix nielseni, sp. nov. (Lepidoptera: Yponomeutoidea: Plutellidae s. l.) Invertebrate Systematics 17: 59–66.

[NC91] Nielsen, E. S., & I. F. B. Common. 1991. Lepidoptera (moths and butterflies). In: CSIRO. The Insects of Australia: A textbook for students and research workers 2nd ed. vol. 2 pp. 817–915. Melbourne University Press: Carlton (Victoria).

[P27] Philpott, A. 1927. The maxillae in the Lepidoptera. Transactions and Proceedings of the New Zealand Institute 57: 721–746.

[P92] Poinar, G. O., Jr. 1992. Life in Amber. Stanford University Press: Stanford.

[P66] Powell, A. W. B. 1966. The molluscan families Speightiidae and Turridae: an evaluation of the valid taxa, both Recent and fossil, with lists of characteristic species. Bulletin of the Auckland Institute and Museum 5: 1–184, pls 1–23.

[RD77] Richards, O. W., & R. G. Davies. 1977. Imms’ General Textbook of Entomology 10th ed. vol. 2. Classification and Biology. Chapman and Hall: London.

[RJ93] Ross, A. J., & E. A. Jarzembowski. 1993. Arthropoda (Hexapoda; Insecta). In: Benton, M. J. (ed.) The Fossil Record 2 pp. 363–426. Chapman & Hall: London.

[VC07] Villanueva, R. T., & C. C. Childers. 2007. Insect predators of eriophyids on Florida citrus including a new predacious Lepidoptera and two cecidomyiids. In: Morales-Malacara, J. B., V. M. Behan-Pelletier, E. Ueckermann, T. M. Pérez, E. G. Estrada-Venegas & M. Badii (eds) Acarology XI: Proceedings of the International Congress pp. 391–395. Instituto de Biología and Faculdad de Ciencias, Universidad Nacional Autónoma de México, Sociedad Latinoamericana de Acarología: México.

[WM66] Wallace, A. R., & F. Moore. 1866. List of lepidopterous insects collected at Takow, Formosa, by Mr. R. Swinhoe. Proceedings of the Zoological Society of London 1866: 355–365.

[WFS04] Winks, C. J., S. V. Fowler & L. A. Smith. 2004. Invertebrate fauna of boneseed, Chrysanthemoides monilifera ssp. monilifera (L.) T. Norl. (Asteraceae: Calenduleae), an invasive weed in New Zealand. New Zealand Entomologist 27: 61–72.

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